Golgi complex

History
L. St. George (1867) discovered certain organelles in cells which were later on called Golgi bodies or
Golgi complex. Camillo Golgi (1898) used silver salts and osmium tetroxide for impregnation of cells by
which presence of lipid material was identified. By using methylene blue stain certain particles were
stained in cytoplasm which were considered as precursor of Golgi bodies.
t was by these staining reactions that the Italian physician Camillo Golgi in 1898 first recognized the
complex in nerve cells of the barn owl and cat. Some workers, like Baker, examined the bodies by using
Sudan black stain and suggested that variations in morphology of these bodies are mainly due to
changes in their secretary activity.
According to Cowdry (1924), Golgi apparatus is an area of cytoplasm, frequently of reticular shape, often
as large as nucleus and sometimes definitely located with regard to cellular polarity. Holmgren called it
as trophospongium (i.e., consisting of a network of fibres packed with secretory granules). Cajal called
it as Golgi-Holmgren body.
Work of Bowen indicated that Golgi apparatus exists in plants in the form of osmiophilic platelets. Thus,
Golgi apparatus has been variously called as Golgi bodies, dictyosomes, lipochondria, internal reticular
apparatus, canalicular system or trophospongium by different workers. In plant cells and the lower
invertebrate cells they are not as prominent as in vertebrate cells.
Occurrence:
Golgi complex is found in all cells except blue- green algae, mycoplasmas and bacteria (i.e., prokaryotic
cells), eukaryotic cells of mature sperm, red blood cells in animals, certain fungi, sperm cells of
bryophytes and Hervdophytes, and sieve tube cells in plants. In plant cells it has been collectively called
the dictyosome and may consist of many flattened sacs.
They are found scattered throughout the cytoplasm. It occupies different positions in different kinds of
cells it is polar, between nucleus and periphery in cells of ectodermal origin; around nucleus in
neurons, and located elsewhere in other cells (Bloom and Fawcett, 1975).
Maintaining a healthy Golgi complex depends upon the presence of a cell nucleus. In the absence of cell
nucleus, Golgi complex decreases in size and may disappear. After the appearance of nucleus in an
Amoeba, small curved cisternae of the Golgi complex reappear.
Starvation reversibly attenuates the Golgi complex, which redevelops soon after the cells are resupplied
with nutrient. This occurs only after the rough ER has redeveloped. Therefore, proteins needed for
reconstituting the Golgi complex must come from the ER (Northcote, 1971).
Terminology of Golgi Material:
Sosa has suggested following nomenclature for Golgi material
1. Golgiokinesis: Division of Golgi apparatus during nuclear division.
2. Golgiosomes: Corpuscles produced by golgiokinesis are called as golgiosomes which are described as
Golgi material in invertebrates.
3. Golgiolysis: Process of dissolution of Golgi apparatus.
4. Golgiorrhexis: Fragmentation of Golgi apparatus.
5. Golgiogenesis: Formation and differentiation of Golgi body during embryology.
6. Golgio-cytoarchitecture: Study of structure of cell in relation to Golgi apparatus.
Shape and size:
Morphology of this complex is variable depending upon the type of cell in which these are present.
In fully mature and functional cells this complex is well developed (e.g., secretory cells and neurons),
where it is clearly a reticulate structure, while in non-functional cells it is poorly developed. In plant cells
Golgi bodies are about 1 to 3 (mu) in length and about 0.5 (mu) in height. These are basically identical
in structure to animal cells Golgi complex.
Structure:
It is still debatable that there is a direct morphological relation between the endoplasmic reticulum and
the Golgi complex. However, it was early recognized by its affinity for osmium of silver-containing stains.
In modern days its existence has been clearly demonstrated with the electron microscope. They are
found in living cells of both animals and plants, immersed in their cytoplasm.
Structurally (electron microscope), fully developed Golgi complex is disc-shaped, having (i) central
flattened plate-like sacs or cistemae, (ii) peripheral interconnecting tubules, (iii) more peripheral
(terminal) vesicles, and larger Golgian vacuoles filled with an amorphous or granular material. Surface of
the Golgi complex is covered by a unit membrane.


Various workers have given various names to this complex, such as dictyosome, idiosome, lipochondria,
Golgi body, Golgi substance, Golgi apparatus and Golgi complex. Generally the name Golgi complex is
used for the material invertebrates and dictyosome in invertebrates and plants. However, the Golgi
complex is not found in bacteria and blue green algae.
The shape of Golgi complex is quite variable in somatic cells of plants and animals. Even in the same cell
there are variations with functional stage. However, the shape is characteristic for each cell type. Their
appearance depends partly on the position of cells within the plant or animal body and partly on the
way in which cells are prepared for microscopic study.
These bodies look like the complexes of the droplets and therefore, termed as Golgi complex. They
appear thin plate like layers of irregular arrangement. Each complex consists of stacks of parallel
lamellae. In some cases, it occurs as a dense reticulum of anastamosing trabeculae while in others as an
irregular plaque, a ring, hollow spheres united together.
In nerve cells it occurs as a reticulum of wide meshes around the nucleus. The size of Golgi complex is
also variable. It is small in muscle cells but quite large in the nerve and gland cells. Generally the
complex is well developed in an active cell while in old cells the complex diminishes in size and
ultimately disappears.

The electron microscopic observations reveal as follows. The Golgi complex comprises of 1. Flattened
sacs or cistemae, 2. Large clear vacuoles and 3. Clusters of dense vesicles.


[I] Flattened sacs or cisternae:
These sacs are flat, elongated and tubiform membrane enclosed structures which are closely packed in
parallel bundles (stack) lying one above the other. Number of sacs per Golgi body or complex varies
from 4 to 7 (Hall et al., 1974) and it may be upto thirty or more in lower organisms.
These sacs consist of a double membranes or lamellae, continuous at the two ends, enclosing a thin
cavity of about 520 A. The membranes are mainly lipoproteinous in nature. Each stack of cisternae
having a proximal or forming face, which is generally convex and a distal or maturing face of concave
shape, which encloses large secretory vesicles. The proximal or forming face is characterized by the
presence of small transition vesicles or tubules that converge upon the Golgi cisternae, forming a kind of
fenestrated plate.
These transition vesicles are thought to form from the endoplasmic reticulum and then migrate to the
Golgi, where they coalesce with each other to form new cisternae. The sacs generally have a narrow
lumen sometimes expanded at its ends.
The contents of lumen have electron density almost equal to that of hyaloplasm, though it is variable
depending on the functional state of Golgi complex. These sacs usually lie in parallel arrays but they may
diverge especially at the periphery of stack.
In exocrine cells of pancreas these sacs are arranged in bundles, each bundle consisting of several
membrane pairs separated from other group by a space of about 130 A. Each membrane is 60-70 A thick
and both the membranes of a sac are separated by a space of 60-90 A wide. In other holocrine and
apocrine cells intervening spaces between sacs may vary from 50 to 200 A wide.
From periphery of cisternae arises an anastomosing flat network of tubules of 300 to 500 A in diameter.
[II] Vesicles or tubules:
These are small drop-like structures smaller in size about 600 A in diameter, and attached with the
tubules or cisternae. These are also derived from tubules by simple budding of sacs or constriction of the
ends of tubules. Thus, liberated particles of sacs are vesicles which become dispersed in the surrounding
hyaloplasm. Two types of vesicles may be connected with the tubules:
1. Smooth vesicles are some what flattened and are of 20 to 80 mu (or 200 to 800 A) in diameter and
contain secretory material. Thus they are often called secretory vesicles. These are located within the
tubular network or sometimes near the centre of the stack of cisternae.
2. Coated or rough vesicles are spherical outgrowths having about 50 mu (or 500 A) diameter. Their
surface is rough. These are found at the ends of the tubules or cisternae.
[III] Vacuoles:
These are large, sac-like globular irregular empty structures situated at the distal ends of sacs. These
vacuoles are formed essentially by enlargement and consequent swelling of sacs, i.e., vacuoles are
swollen sacs in which vacuolar space becomes widened. In limpet, Patella vulgata, large vacuoles vary
from about 760 A to about 6000 A in diameter, whereas in the convoluted tubules, vacuoles range from
500-2000 A.
Intercisternal elements, 70 to 80 A in diameter, may be found between adjacent cisternae. They appear
to be fibrous elements and have no known function.
Kirkman and Severinghaus state that the Golgi body consists of a fibrous network, ring or cylinder, a very
irregular fenestrated plate and collection of small spheres, etc. This is usually well developed in the
stage of cytomorphosis and tends to decrease in size as cells become older. In gland cells the structure is
fairly developed. Pollister says that Golgi body appears as a plate-work of irregular shape and of uniform
thickness.
Generally in vertebrate somatic cells, the Golgi body consists of a dense network situated near the
nucleus. In germ cells it is in the form of rods or granules.
On its staining properties, Golgi body consists of two parts, an outer which absorbs osmium and silver
(argentophilic) and inner which is osmic or argentophobic (Richardson, Bourne). Baker (1944) found that
the Golgi body is composed of four parts
(1) Neutral red vacuoles
(2) Dense lipoid-containing substance
(3) Diffuse lipoid-containing substance
(4) Golgi product
Neutral red vacuoles in Golgi body have been shown by Hibbard in digestive tract cells of chicks.
Main functions of Golgi complex are as follows:
Golgi apparatus appears to play an important role in the storage, packaging and secretion of certain cell
products. It is involved in the formation of lysosomes and other enzyme-containing cellular inclusions,
and in the formation of secretory granules in cells such as those found in the pancreas, pituitary and
mammary glands, and mucous-secreting glands of the intestine and in many other cell types.
Functions:
In general Golgi complex is of vital importance and serves many functions:
1. Absorption of compounds:
Hirsch et al., have discovered that, when iron sugar is fed to an animal, iron becomes absorbed on Golgi
bodies (Kedrowsky). Van Teel has shown that Golgi systems also absorb compounds of Cu and Au (gold).
Kedrowsky has shown that Golgi bodies of Opalina can absorb bismutose (compound of albumin and
bismuth) and protargol (compound of albumin and silver). Palay and Kartin (1956) have indicated that
Golgi complex is concerned with the storage and absorption of lipids.
Thus, Kirkman and Severinghaus state that Golgi apparatus acts as a condensation membrane for the
concentration of products produced elsewhere into droplets or granules by losing water, which are
transported to the cell surface for export.
These products may be lipids, yolk, bile compounds, enzymes and hormones, etc. The proximity of Golgi
and contractile vacuoles in protozoa may confirm the concentration behaviour of Golgi membranes.
Golgi membranes may remove water from the products of synthesis during the formation of secretory
granules.
2. Formation of secretory vesicles and secretion:
The principal function of Golgi complex is secretion. In several types of cells, synthetic products from the
rough endoplasmic reticulum are transferred to Golgi region, from where they are liberated from the
cell through plasma membrane by pinocytosis. Secretory function of Golgi seems to be well founded
experimentally.
3. Helps in enzyme formation:
Bowen speaks of Golgi apparatus as a great intracellular centre of enzyme formation. Moricard has
shown that Golgi body helps in the production of follicular fluid from granulosa cells of ovary.
It may release zymogen granules (inactivated pancreatic enzymes) which arise from cisternae into
secretory vesicles. Secretory vesicles containing granules migrate to cell surface where the membrane of
the secretory vesicle and the cell membrane merge, releasing the contents into pancreatic ducts from
where they pass into intestine as active digestive enzymes.
4. Production of hormones:
Golgi body in endocrine cells helps in secretion of hormones. Cowdry has suggested that any harm to
Golgi apparatus in thyroid gland cells will result in decline in secretion of its hormone.
5. Storage of protein:
Vacuoles and vesicles which are the main components of Golgi complex become filled with protein-
lipoid material for storage. These stored products help in secretory action.
6. Formation of acrosome:
It forms the acrosome of sperm during sperm maturation as shown in Fig. 2. The cisternae of Golgi
complex are arranged in a cup-shaped pattern, the lamellae stacked in parallel. From the periphery of
lamellae, small vesicles or vacuoles are pinched off.
Gradually the system of cisternae is replaced by more vesicles and tubules and within some of these
vesicles small granules appear. These granules represent secretory products within Golgi complex. Some
of the granule-containing vesicles coalesce to form a single acrosome within a large vesicle, which
comes to lie on the surface of the sperm nucleus.

The vesicle gradually spreads over more of the nuclear surface and eventually collapses on the nuclear
membrane to form the cap material, acrosome.
7. Formation of intracellular crystals:
In the marine isopod, Limnaria ligmorum, which is a burrowing form, there are present midglands whose
cells consist of crystals. These range up to 30 in length and 15(1 thick. It has been proved that these
crystals are formed by Golgi complex and are known to contain protein and iron. They are without
enclosing membrane and usually spheroidal in shape. They are concerned with the secretory activity.
8. Milk protein droplet formation:
In lactating mammary gland of mice are produced protein droplets which are related with Golgi
complex. These droplets usually open on to cell surface by fusion of their enclosing membrane with
plasma membrane.
9. Formation of plant cell wall:
Golgi bodies of plant cells synthesize all polysaccharides such as pectin, hemicellulose and microfibrils of
a-cellulose. These are packaged in vesicles for secretion. For example, pectin and other mucilaginous
substances of the plant cell wall are synthesized in the Golgi and are packaged in vesicles for secretion.
Golgi bodies during mitotic cell division form a cell plate at the centre of spindle. This cell plate is
gradually enlarged and thickened by deposition of pectic substances, hemicellulose and micro fibrils of
a-cellulose secreted by Golgi bodies.
10. Glycoproteins secretion:
Glycoproteins are formed in the Golgi complex by the attachment of carbohydrate to the protein
products of the endoplasmic reticulum.
After enzymes are synthesized and accumulated in vesicles of endoplasmic reticulum, the vesicles
migrate to the Golgi complex, where they fuse with the Golgi membranes. In the Golgi complex, the
newly synthesized products of ER are concentrated, and some of the proteins are modified by the
addition of carbohydrates or other prosthetic groups.
The concentrated products then accumulate in the edges of Golgi cisternae, where they are packed into
small vesicles that bud off from the edges of cisternae and are released into the cytoplasm. Here smaller
vesicles may fuse into larger ones. If the vesicles contain secretory product, they will migrate to the
plasma membrane, where they release their products into the surrounding environment by exocytosis.